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Female Ectodermal Genitalia of the Taxa of the Kelisia Guttula

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Female Ectodermal Genitalia of the Taxa of the Kelisia Guttula Female ectodermal genitalia of the taxa of theKelisia guttula - group (Homoptera Fulgoromorpha Delphacidae): not only an example for specialized coorganization with male genitalia without obvious reasons, but also a character set apt for species discrimination in westpalaearctic taxa? R e in h a r d R e m a n e & Ad a l g is a G u g l ie l m in o Keywords: Kelisiinae, evolutionary coorganization, “hypertrophic” specialisation, “orthogenetic evolution”, biogeography (Spain, France, Central Europe), taxonomy, Kelisia occirrega n. sp. Abstract: In the course of evolution male and female external and internal ectodermal genitalic structures in the taxa of the monophyletic Kelisia gwffw/a-group have been enlarged and differentiated in a very special way in comparison to plesiomoiphic taxa of Kelisiinae. Functional reasons for this “orthogenetic” evolution are not yet known, as it seems. In evolutory “coorganization” the evolution of a proportionally very large genital segment and an extremely long, thin and flexible aedeagus in the males is “answered” (for there seem to be - other than in some other insect taxa, e. i. Diptera Tephritidae - no special “female” functions like oviposition or “sperm handling” which may act as “selective forces” on these structures, by this changing their morphology) by the females with the development of a special duct of approximately “edeagal length” guiding and keeping the male aedeagus during copulation. This “edeagal duct”, which morphologically is “folded off’ from the genital room, starts at the dorsal base of the female’s genital room and ends opposite the depart of the oviductus communis (leading to the spermatheca as well as to the ovaries). It is much longer than the genital room, but it remains connected with it by a thin ligament. It is provided with a variety of sclerotized structures. This could be interpreted as a “lock and key” mechanism to prevent successful mating between specimens belonging to different species, but such matings are prohibited already by species-specific vibratory signals acting as “premating isolating mechanism”. As in the K. guttula-group there exist several species-groups containing externally very similar species, in which the females could not be identified down to species up to now (resulting in “handicaps” for biogeographical and ecological analyses), we examined these structures looking for species-specific characters enabling the safe identification not only of males, but of females, too. As a result all females of this group now may be safely identified due to such species-specific characters present in their internal ectodermal genitalia. The populations of “Kelisia irregulata Haupt” from Spain and western France have shown to be that different from the allopatrically distributed central european populations, that they seem to be a genpool of their own. So we decided to describe these populations as a species of its own: Kelisia occirrega n. sp. (Type locality: Spain, Prov. Pamplona, Isaba in the Valle del Roncal in the southwestern Pyrenees). This taxon ought to keep its species rank unless intermediate, transitional specimens will be discovered in the “gap” between the two taxa (situated in eastern France) indicating the existence of a “hybrid belt”. As known, one of the aims of taxonomic research is to detect character states, which enable a researcher safely to decide to which of two or more otherwise very similar species a certain specimen belongs. Most favourable are those character states, which may be easily seen and persist even if the specimen is dead and conserved by appropriate techniques. Unfortunately, in many groups of animals this aim has not yet been achieved: one of the sexes and/or juvenile specimens have to remain unidentified. This situation exists also in the “guttula-group” of the planthopper genus Kelisia Fieber. The number of species recognized in this species-group has not only increased during the past years, but the characters used for species recognition and discrimination have changed, too: while in former times (R ib a u t 1934; W a g n e r 1939) it was possible to recognize the alleged “species” not only by characters of the male genitalia but also by external characters like size, shape, proportions, colour and markings, more recently species have been established or described (R e m a n e & Ju n g 1995, G u g l ie l m in o & R e m a n e 2002) which could be discriminated from their nearest relatives no longer by external characters but by the structure of the male genitalia only. In most of these cases their females had to remain unidentified. As such a situation is unfavourable for ecological as well as for biogeographical research, we started an examination of the internal ectodermal parts of the female genitalia in order to find out, whether there might exist species-specific characters or not. In the monophyletic “ Kelisia gwtfw/a-group” (consisting up to now of 10-11 species: 3 in the eastern part of the Palaearctic Region only: xiphura Vilbaste, bispinifera Dlabola and asahinai Hori, and 8 confined to or reaching into the western Palaearctic: guttula Germar sensu Wagner, sima Ribaut, italica Guglielmino & Remane, irregulata Haupt, vittipennis J. Sahlberg, haupti Wagner, halpina Remane & Jung, hagemini Remane & Jung) the internal female genitalia are very specially constructed1. Directly at the dorsocephal side of the genital-opening following the “Atrium-Platte” there is developed a special duct accompanying the basal part of the “Genitalraum”, i. e. the part extending between the cephalodorsal basis of the GVIII and the origin of the duct leading to the ovaries and the receptaculum seminis. This special duct “directs” and “keeps” the male aedeagus during copulation. To succeed in this, it is provided with more or less extended sclerotized walls: some of these sclerotizations are located on its dorsal side, and one at the "end” directly before this duct enters again the genital room. The male of the taxa of the K guttula-group possess not only an absolutely and relatively to the size of the body very enlarged genital segment but also a thin, flexible and extremely elongated aedeagus (see amongst others W a g n e r 1963 and Asc h e 1985), which at rest lies curved in a semicircle in the distal half of the male abdomen. “Coorganized” with the length of the aedeagus, the duct in the female genitalia (but not the corresponding part of the genital room!) is considerably elongated and by this is forced to “depart” from the genital room, forming in some taxa a large, undulated, asymmetrically situated lobe. This lobe always stays connected over its complete length with the genital room by a thin, membraneous ligament. By this a kind of separation 1 The non-westpalaearctic taxa K. xiphura Vilbaste, K. bispinifera Dlabola and K. asahinai Hori, 1982 have not yet been investigated due to the lack o f material. The description oK. f asahinai Hori unfortunately lacks important details of the structure o f the male genitalia (only tip of the aedeagus shown, subanal appendages neither figured nor described) which render the placement of this species somewhat doubtful. between “copulation duct” and “oviduct” results, which reminds the ditrysic constructions found in other taxa (e.g. Stenocraninae). This character complex - thin, long and curved aedeagus and detached special “edeagal duct” in the females - is a special feature of the taxa of the “ K gwtfw/a-group” not present in any other palaearctic or nearctic groups of Kelisiinae. It may be considered as a special synapomorphy “proving” the monophyly of this group of taxa2. To observe the aforementioned characters, it is necessary to macerate the female abdomen in K(OH)/potassium hydroxid (ca. 10%) and after that using a binocular microscope to wash and dissect it in water. After that it may be transferred either as usual into glycerol for detailed observations or (if parts like genital room, oviduct, and bursa copulatrix are to be kept inflated) in water to which formaldehyd has been added (ca. 1:15, see Asche 1985: 27). To examine the details, a magnification of ca. lOOx is recommended. The results of our investigations have to be considered preliminary, for from many species we have examined a rather small number of specimens only: the range of variation of these characters within a population as well as within the geographical range of each taxon thus might be larger than shown here and than known up to now. Nevertheless, the results obtained so far seemed that promising with respect to the aim of species discrimination in this K. guttula-gcoup, that we decided to publish them already now: so these our results presented here may be corroborated, or corrected by other homopterologist. The characters, in which we think to have found differences between the species of this group mainly are: 1st The absolute and relative length of the edeagal duct. This length apparently (and as expected) is ± correlated with the length of the aedeagus. But exact measuring is difficult (and time-consuming), as this duct is bent and twisted in all directions especially in those taxa, in which it is very long. In addition the way in which this duct is bent differs between specimens of the same species (even if these specimens were caught at the same data in the same place). Nevertheless - as shown by the figures - there is a very short duct in K. hagemini Remane & Jung (Fig. 1), this duct is distinctly longer in K. haupti Wagner (Fig. 2) and much longer in K. halpina Remane & Jung (Fig. 3). These three species were separated in 1995 only (R emane & Jung ) by morphometric examination of their male genitalia, they are very similar in size, proportions and markings and at that time their females could not be reliably separated.
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